Western Indian Ocean J Mar. Sci. Vol. 10, No. 2, pp. 137-154, 2012 © 2012 WIOMSA 243808 The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa Jennifer M. Olbers 1 and Yves Samyn 2 1 Zoology Department, University of Cape Town, Private Bag X3, Rondebosch, 7701, South Africa; 2Belgian Focal Point to the Global Taxonomy Initiative, Royal Belgian Institute of Natural Sciences, Belgium. Keywords: Ophiocoma, neotype, taxonomy, Ophiuroidea, Indo-West Pacific, Western Indian Ocean, KwaZulu-Natal. Abstract-This study raises the number of Ophiocoma species recorded in South Africa from four to eight. All species are briefly discussed in terms of taxonomy, geographic distribution and ecology. In addition, the juvenile of 0. brevipes, found on the underside of adult Ophiocoma brevipes specimens, is described in detail. A neotype is designated for 0. scolopendrina. INTRODUCTION The circumtropical family Ophiocomidae holds some of the more dominant and conspicuous ophiuroid species present on coral and rocky reefs. The family is rich, with eight genera, two of which were relatively recently reviewed by Devaney (1968; 1970; 1978). One of these, Ophiocoma Agassiz, 1836, is well represented in the tropical to subtropical waters of KwaZulu-Natal in South Africa and its constituent species are documented here. Ophiocoma species are difficult to identify since some of the distinctive taxonomic characters, such as the shape of the oral and dental plates and their associated papillae (see Devaney 1970), are not easily assessed or can change during growth, as is also the case for the number of arm spines, the disc armament and the size of the dorsal arm plates (Sumida et al. 1998; StOhr 2005; StOhr et al. 2008). In addition, some species can change their colouration from day to night (Hendler 1984). Corresponding author: JMO E-mail: [email protected] The Indo-Pacific distribution of Ophiocoma has been dealt with by several authors (e.g. Clark & Rowe 1971; Cherbonnier & Guille 1978; Rowe & Gates 1995). Clark and Rowe (1971) listed 11 species from the Indo-West Pacific (including the Red Sea and the Persian Gulf). Since then, a few new species have been added (Rowe & Pawson 1977; Bussarawit & Rowe 1985; Soliman 1991; Benavides-Serrato & O'Hara 2008), bringing the total number of valid species in the Indo-Pacific to 13 1• The ophiuroid fauna of southern Africa have been studied by many specialists (Bell 1905; 1909; Clark 1923; Mortensen 1925; 1933; Balinsky 1957; 1969; Clark 1974; 1977; 1980; Clark & Courtman1 Ophiocoma anaglyptica Ely; 0. aegyptiaca Soliman, 1991, 0. brevipes Peters, 0. cynthiae BennavidesSerrato & O'Hara, 0. dentata Muller & Troschel, 0. endeani Rowe & Pawson, 0. erinaceus Muller & Troschel, 0. occidentalis Clark, 0. pica Milller & Troschel, 0. pus ilia (Brock), 0. schoenleinii Milller & Troschel, 0. scolopendrina (Lamarck), 0. valenciae Muller & Troschel. JENNIFER M. OLBERS and YVES SAMYN 138 Stock 1976) but, since Clark and CourtrnanStock's (1976) work, no Ophiocoma species have been added to the fauna of southemAfrica. Samynand Thandar (2003), however, mention that de novo sampling along the coast of KwaZulu-Natal revealed new ophiuroid many records. Unfortunately, the latter authors did not include their species list. Until now, only four species of Ophiocoma have been recorded in South Africa (Clark & Courtman-Stock 1976): 0. erinaceus Muller & Troschel 1842, 0. pica Muller & Troschell842, 0. scolopendrina (Lamarck 1816) and 0. valenciae Muller & Troschel 1842. Recent sampling along the north-east coast of South Africa (see also Samyn & Thandar 2003), yielded four additional Ophiocoma species. Moreover, we discovered an undocumented association between individuals which we believe to be adult and juvenile 0. brevipes. Abbreviations: A.L. Ann length Disc diameter KwaZulu-Natal KZN MHNG Museum d 'Historie Naturelle, Geneve, Switzerland MNHN Museum national d'Historie Naturelle, Paris, France RMCA Royal Museum for Central Africa, Tervuren, Belgium Natural History Museum, London, United Kingdom NHM iZiko South African Museum, Cape Town, South Africa SAM D.D. ZMB Museum fiir Naturlrunde an der Universitat Humboldt m Berlin, Berlin, Germany MATERIALS and METHODS Specimens were collected by hand in the intertidal zone and by SCUBA diving up to 32 m depth, during six expeditions (August 1999, July 2000, February 2001, July 2003, January 2010 and October 2010) at several localities along the coast ofKZN, South Africa (Fig. 1). Specimens were anaesthetized by placing them in freshwater and gently manipulating them so that their arms were extended as far as possible until they ceased moving. Thereafter, they were put in 100 %buffered alcohol for one day and transferred to 70 % buffered alcohol for transport to the laboratory where they were dried for permanent storage. The specimens were photographed after preservation. Specimens are deposited in the collections of the RMCA(Belgium) and in the SAM (South Africa). These collections were examined together with other vouchers in the RMCA (e.g. from Kenya, the Seychelles and Inhaca; see also Clark 1980; 1984) as well as some ophiocomids from the Indo-Pacific Ocean in the MNHN (France). "'I" "1". a:~ •I" ~ "1" "'f' "'I" , "1" ... "1' -~. ,.,. "f' ...... KOSI Bay Bhan ga Nek ...,.. .,.,,. ,..,.. .,.... . ., Umhlalt Durban /"' .,,.. AltWal Shoal ar1<R)'Tl., 7no ~'-.. •.!.,..,_,-.; r.. ~$ ortSt Johns Umgazana olora ..:.. ,... ,.,. ,;, 90 180 ...:.. """' , ... Figure 1. Sites where Ophiocoma species have been recorded in South Africa. 270 li'.O ,..,., 360 Kllomoters .... ,.:.., ,..,. The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa 139 Table 1. Records of the present South African collection of Ophiocoma species, showing location, depth, museum collection numbers and the number of specimens. Valid species name Location Ophiocoma brevipes Aliwal Shoal Sodwana Bay 20 14 Sodwana Bay (2 mile reef) 15 15 11 Sodwana Bay (Mbibi) Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Sodwana Bay (9 mile reef) Sodwana Bay (9 mile reef) Sodwana Bay (2 mile reef) Sodwana Bay (Mabibi) Aliwal Shoal Sodwana Bay (Diep Gat) Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Aliwal Shoal Sodwana Bay Bhanga Nek Aliwal Shoal Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Depth (m) 11 6 15 16 13 10 11 15 14 13 Collection number (no. of specimens) RMCAMT2199(1) RMCAMT 2207 (1) RMCAMT2175 (1) RMCAMT2215(1) RMCAMT2191 (1) RMCA MT 2157 (1) RMCA MT2192 (1) RMCA MT 2225 (1) RMCAMT2222 (1) RMCAMT2194 (1) RMCAMT2195 (1) RMCAMT2198(2) RMCAMT2179(1) RMCAMT2212(1) RMCAMT2193 (1) RMCA MT2148 (3) RMCA MT 2208 (2) RMCA MT 2203B1S (2) RMCAMT2!99 (I) SAMA28111 (1) RMCA MT2341 (2, Juvenile) SAM A28112 (2, Juvenile) Sodwana Bay (2 mile reef) 16 20 14 15 14 Ophiocoma cf dental a Sodwana Bay 20 RMCAMT2380(1) Ophiocoma doederleini Bhanga Nek (Saxon Reef) Sodwana Bay (2 mile reef) Bhanga Nek 20 15 16 RMCA MT 2249 (1) RMCAMT2250(1) RMCA MT 2203B1S (2) Ophiocoma erinaceus Sodwana Bay (5 mile reef) Bhanga Nek (Saxon Reef) Sodwana Bay (2 mile reef) Bhanga Nek Sodwana Bay (1/4 mile reef) Sodwana Bay (7 mile reef) 18 20 15 RMCAMT2136(3) RMCAMT2142 (2) RMCAMT2253 (1) RMCA MT 2155 (!,juvenile) RMCAMT 2220 (!,juvenile) RMCAMT2349(1) Ophiocoma pica Qolora, Eastern Cape Ophiocoma pus ilia Aliwal Shoal Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Bhanga Nek Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Aliwal Shoal Sodwana Bay (Deep sponge) Aliwal Shoal Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Sodwana Bay (2 mile reef) Ophiocoma scolopendrina Umgazana, Eastern Cape Ophiocoma valenciae ParkRynie Park Rynie Sodwana Bay (5 mile reef) Sodwana Bay (5 mile reef) Sodwana Bay (5 mile reef) Aliwal Shoal Aliwal Shoal Aliwal Shoal Aliwal Shoal Aliwal Shoal Durban Durban Port St Johns Umhlali 20 10 17 Intertidal 20 13 l3 15 20 15 14 16 32 13 10 10 14 SAM A 23248 (2) RMCA MT2345 (1) RMCA MT2219 (I) RMCAMT2200 (I) RMCAMT2346 (I) RMCAMT2337 (I) RMCAMT2217 (1) RMCA MT 2214 (I) RMCA MT 2218 (I) RMCAMT2380(1) RMCA MT 2153 (1) RMCAMT2221 (1) RMCAMT2149(1) RMCAMT2381 (I) SAM A 23093 (2) Intertidal Intertidal 18 16 18 16 13 13 13 13 Intertidal Intertidal Intertidal Intertidal RMCA MT 1754 (3) RMCAMT1755(1) RMCAMT 1748 (1) RMCAMT1749(1) RMCA MT 1750 (3) RMCA MT 1756 (I) RMCA MT 1751 (1) RMCAMT 1753 (I) RMCAMT 1752 (I) RMCAMT 1747 (1) SAM A 23250 (1) SAM UCTECOLCOLL. D 178(1) SAM MEIRING NAUDECOLL. J 9_1 (6) SAM UCT ECOL COLL. U 23A (I) 140 JENNIFER M. OLBERS and YVES SAMYN RESULTS Details of the collection of different Ophiocoma species sampled along the KZN coast of South Africa are presented in Table l. SYSTEMATIC ACCOUNT We provide a taxonomic description and detailed geographical distribution for Ophiocoma species that are new records for South Africa. For the other well-known species, we only provide a summary and cite major works which provide detailed information on their taxonomy and distribution. Order Ophiurida Miiller & Troschel, 1840 Family Ophiocomidae Ljungman, 1867 Subfamily Ophiocominae Matsumoto, 1915 Genus Ophiocoma Agassiz, 1836 (Type species Ophiura echinata Lamarck, 1816 (by subsequent designation ofClark 1915)). When distal lobe angular, proximal end usually more or less truncated to match interradial separation of adoral shields. Dorsal arm plates wider than long, fan-shaped, oval or hexagonal. Ventral arm plates more or less square-shaped, proximal side straight, distal side straight to concave. Two tentacle scales, at least in first arm segments, sometimes only one towards tip. Ophiocoma brevipes Peters, 1851 (Plate 1A-G, 2A-C) Ophiocoma brevipes Peters 1851: 466; Marktanner-Turneretscher 1887: 303; de Loriol 1893: 25, 26, pl. 23, Fig. 4; Clark 1908: 296; Clark 1911 : 256; Koehler 1922: 319-322, pl. 72, Diagnosis (after Devaney 1970: 9; Clark & Courtman-Stock 1976: 172-173). Majority of species large with D.D. often exceeding 20 mm; three to seven generally smooth, stout arm spines, sometimes alternating three and four on successive arm segments or on opposite sides of same segment; lower arm spines sometimes flattened and spatulate while upper ones cylindrical or cigar-shaped. Disc generally covered with granules, occasionally concealing scaling on disc and sometimes extending into ventral interradial regions. Oral shields without granules or spines, triangular with three to four, rarely five, contiguous oral papillae, outer one usually widest. Tooth papillae always present, few to numerous, with superficial ones in series with oral papillae. Oral shields large, oval, hexagonal or pentagonal. Plate 1. Dorsal (A) and ventral (B) views of Ophiocoma brevipes; position of juvenile 0. brevipes within the bursal slit of an adult 0. brevipes (C & D); dorsal view of juvenile 0. brevipes (E); ventral view of juvenile 0. brevipes (F); dental plates (internal and external view) of adult 0. brevipes (G). The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa 141 Material examined We examined over 150 individuals of 0. brevipes from South Africa and Madagascar. On seven of them, one to three smaller individuals were attached in such a way that they had their arms slotted in the bursal slit. We have here tentatively treated them as juvenile 0. brevipes. Adults D.D.=5.6-25.2 mm; D.D./ A.L. from 1/3.2 to 1/4.8. Disc colour patterns variable with a combination of light greens, whites, yellows and browns in blotchy star or simply no particular pattern. Disc with small, fine, spherical granules closely packed on both dorsal and ventral side. Oral shields with darker markings, but with no apparent pattern. Dental plate (Plate 1G) between 1.9 and 2.1 times longer than wide, with a wide vertical septum between each oval, slightly elongated tooth foramen, dental papillae region Plate 2. Dental plates of juvenile Ophiocoma brevipes (A); arm limited to approximately 27% spines of juvenile 0. brevipes (B); distal tip of arms of juvenile 0. of dental plate length. Genital brevipes (C); dorsal view of Ophiocoma cf. dentata (D); ventral slits clearly visible, elongated view of 0 . cf. dentata (E); dorsal view of Ophiocoma doederleini and bordered with slightly more (F); ventral view of 0. doederleini (G) .. prominent granules. Arms appear banded (darker) on the dorsal side, some Figs. 6-9; Devaney 1968: 45; Devaney 1970: specimens with light dots or specks on ventral 13; Clark & Rowe 1971: 86, 119; Devaney side of each arm plate along the length of 1974: 151-152; Cherbonnier & Guille 1978: arm. Ventral arm plates nearly as wide as 168-169, pl. X, Figs. 3,4; Sloan et al. 1979: 104; long, bluntly pointed on the proximal side. Clark 1980: 534; Tortonese 1980: 125, Fig. 11; Dorsal arm plates fan-shaped, much wider Humphreys 1981 : 10, 23; James 1982: 39-40, than long. Upper-most spines thickest on the pl. IlB; Clark 1984: 100; Rowe & Gates 1995: proximal part of the arm. Longest arm spine 385; Rowe & Richmond 2004: 3292. less than or equal to the breadth of the dorsal arm plate. Two tentacle scales. Location and status of types - ZMB 961 (1 syntype), ZMB 962 (1 syntype), ZMB 963 (1 Juveniles syntype), ZMB 4660 (3 syntypes). D.D. = 1.5-4.3 mm; D.D./A.L. from 1/1.8 to Type locality - Coast of Mozambique (ZMB 1/2.8. Disc colour brown-grey, both dorsally 961-963); Quirimba Island, Mozambique (Plate lE) and ventrally (Plate 1F). Dorsal (ZMB 4660). side of disc marked with five radial pairs 142 JENNIFER M. OLBERS and YVES SAMYN of bowed brown lines extending from the margin, passing through radial shields and almost reaching centre of disc (Plate lE). One specimen has a faint white dot in the centre of its disc. Radial shields oval, exposed, minute in size, hardly visible and similar in colour to disc. Disc circular in outline and moderately convex, covered dorsally and ventrally by imbricated scales of different sizes and shapes. Disc scales hardly visible dorsally but obvious ventrally. Disc granulation absent on ventral and dorsal surfaces. Oral shields and adoral shields uniformly brownish. Oral shields pearshaped, sometimes with a thickened distal side, hardly longer than broad. Adoral shields triangular, almost touching proximally. Jaws triangular with three to four hyaline-tipped, somewhat pointed oral papillae on each oral plate. Dental plate with one to two teeth and three to four hyaline-tipped dental papillae; lower-most two to three forming a cluster at the apex of the jaw angle (Plate IF). Dental plate (Plate 2A) 1.6 times longer than wide, with a thin vertical septum between each oval, slightly elongated tooth foramen, dental papillae region limited to approximately 28% of dental plate length. Genital slits unarmed, almost reaching the margin of disc. Dorsal arm plates uniformly brown and fan-shaped, first two to four plates contiguous, thereafter well-separated. Ventral arm plates uniform brownish-grey. First two ventral arm plates slightly broader than long with a straight distal edge; thereafter plates gradually become longer, almost rectangular in shape with the distal edge gradually becoming more concave. Arm spines light brown with dark longitudinal line dorsally (Plate 2B). Arm spines longer than length of arm plates, two on first segment, three on segments two to five, thereafter alternating two and three spines on segments six to nine, and two on further distal segments, hyaline and ragged structure. Distally, arm spines with multifid claw-like hooks (Plate 2C). Single tentacle scale over whole arm length apart from largest juvenile (D.D. = 4.3 mm) with two tentacle scales on first two segments. Geographical distribution Tropical Indo-West Pacific region but absent in the Red Sea, the Persian Gulf and north-western parts of the Indian Ocean (Rowe & Richmond 2004). Prior to this study, the only record from southern Africa was Mozambique (Quirimba Archipelago) (Clark & Rowe 1971). Thus the present record extends range to Aliwal Shoal, some 60 km south of Durban, South Africa. Juveniles were found on adults from South Africa, Madagascar and Isles Glorieuses. Ecology Ophiocoma brevipes is associated with coral heads or boulders, on fine to coarse sand and at the base of algal plants in the sandy littoral zone (between 0-54 m depth, cf. Lane et al. 2000). An overview of known microhabitats of 0. brevipes is presented in Sloan et al. (1979). On one living adult, the juvenile was observed to partially move into the genital slit of the adult after collection (Plate IC, D). Remarks The complete absence of granulation on the disc and the loose meshwork structure of the dental plate indicate that the specimens attached to the adult 0. brevipes are most probably juveniles. The colour pattern of these juveniles somewhat resembles a juvenile Ophiomastix venosa which is known to attach itself to Ophiocoma scolopendrina (Four'gon et al. 2007). However, based on the descriptions by Fourgon et al. (2007) and Cherbonnier and Guille (1978), it can be ruled out that the nine specimens studied here are juvenile Ophiomastix venosa, because (i) arm spines of 0. venosa are distinctly longer, are glassy and thin; (ii) jaws of 0. venosa are much more elongated and (iii) ventral arm plates of 0. venosa taper more proximally. The proximally extended adoral shields which nearly meet around the oral shield are indicative of Clarkcoma Devaney, 1970 and the alternating number of arm spines, with the uppermost spine enlarged (approximately The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa two segments long) suggests an affinity with Devaney's (1970) Scolopendrina Group. On the other hand, the appressed morphology of the dental plates is indicative ofDevaney's (1970) Brevipes Group and, therefore, we believe the juveniles are 0. brevipes. If this identification can be confirmed with complementary studies, such as molecular systematics, this would present a new case of parental care. The tips of the arms in the juveniles were found to bear hooks that may assist in gripping (see Plate 2C). In addition to the hooks, the spines have a ragged structure that could also assist in clinging to the adult (see Plate 2B). Hooked spines have been also been observed in ophiuroid species with an epizoic lifestyle (Hyman 1955). Ophiocoma cf. dentata Miiller & Troschel, 1842 (Plate 2D, E) Ophiocoma dentata Muller & Troschel1842 : 99, pl. VII, Figs. 3, 3a; Devaney 1968: 45; Devaney 1970: 13; Clark & Rowe 1971: 86, 119, pl 18, Figs. 2-3; Cherbonnier & Guille 1978: 168, pl. C, Figs. 3, 4; Tortonese 1980: 125, Figs.llA, B; James 1982: 40, pl. IIC, D; Rowe & Gates 1995: 386; Price & Rowe 1996: 76; Rowe & Richmond 2004: 3292. Ophiocoma insularia Lyman 1861: 80-81; Macnae & Kalk 1958: 130. Location and status of types - ZMB 931 (holotype, fixed by monotypy ). Type locality -Unknown according to Muller and Troschel (1842 : 99). However, the ZMB has only one specimen of Ophiocoma dentata in its collection. The catalogue indicates that this specimen was deposited by Deppe, just as is indicated by Muller and Troschel (1842), and that it comes from 'Celebes?' 2, currently known as the Islands of Sulawesi (Indonesia). Material examined D.D. = 14.3 mm, variegated with brown, white and beige, both dorsally and ventrally with the presence of small dark brown spots. Oral 2 MUller & Troschell842 give no indication that they had more than one specimen before them, hence ICZN Art 73.2.2 applies. 143 shields round, as long as wide, with marbled pattern. Dental papillae broad, not extending far into mouth. Dorsal arm plates beige to brown with a whitish-grey patch surrounded by dark brown border on the median distal side, broad and elliptical. Lateral arm plates lighter with several spots. Ventral arm plates light with same spots; sometimes a dark coloured patch is present centrally, square with rounded corners, as wide as long. Arm spines white to beige, broadly and irregularly banded once or twice with light brown. Upper arm spines thick, blunt, somewhat flattened and slightly shorter than the lower ones. Tentacle scales, two. Ecology According to Devaney (1970), this species frequents the sub-littoral zone, under boulders or associated with coral and coral debris on a sand or rubble substratum. Geographical distribution Ophiocoma dentata has a tropical, Indo-West Pacific distribution (with the exception of the Red Sea and north-western Indian Ocean (Rowe & Gates 1995; Rowe & Richmond 2004). Prior to this study the most southern records originated from Inhaca Island (Mozambique) (Macnae & Kalk 1958; as 0. insularia Lyman 1862). Remarks This specimen, at first examination, bears resemblance to Ophiocoma brevipes. Devaney (1970) pointed out three means by which 0. brevipes can be separated from 0. dentata (and 0. doederleini; see hereunder) for specimens of similar size. First, by comparing the arm spine sequence: while 0. brevipes presents five arm spines on segments four to seven, five to six arm spines up till segment 11, and two to three arm spines thereafter, 0. dentata has only four (occasionally five) arm spines on segments four to seven, four arm spines up until segment 13 and three arm spines thereafter. Secondly, in 0. brevipes the longest arm spine rarely exceeds the breadth of the ventral arm plate, while in 0. dentata (and 0. doederleini) the longest arm spine greatly exceeds the ventral arm plate. Thirdly, 144 JENNIFER M. OLBERS and YVES SAMYN by comparing the pigmentation: while 0. brevipes has a uniform white or cream colour on the oral side of arms and the oral plates and shields, the other two species have a more grey, brown or variegated colouration. However, our specimen differs somewhat from the typical Ophiocoma dentata as reported by Devaney (1970). First, the arm spine sequence of our specimen has three arm spines on its four first segments, segments five to fifteen have four arm spines, and thereafter three arm spines, which differs from the arms of 0. dentata where we never observed five arm spines on segment seven. Second, the colouration of the dorsal arm plates is beige to brown with a whitish-grey patch that is bordered by dark brown on the median distal side, which differs from the description given by Devaney (1970, Fig. 21 ). Finally, the white to beige arm spines are broadly and irregularly annulated with light brown. Given these three differences, we are reluctant to describe the single specimen as a new species until more material becomes available. Ophiocoma doederleini de Lorio1, 1899 (Plate 2F, G and Plate 3A, B) Ophiocoma doederleini de Loriol 1899: 30, pi. 3, Fig. 2; Devaney 1968: 69; Devaney 1970: 1218, Figs. 18, 14, 22, Table 2B, 3; Devaney 1974: 154; Sloan et al. 1979: 104, Figs. 8-10; Clark 1980: 534; Humphreys 1981: 10, 24; Clark 1984: 100; Rowe & Gates 1995: 396. Plate 3. Dorsal views of reticulated (A) and spotted (B) forms of Ophiocoma doederleini; dorsal view of Ophiocoma erinaceus (C); dorsal view of juvenile 0. erinaceus (D); ventral (E) and dorsal (F) views of Ophiocoma pica (from Mahe, Seychelles). Ophiocoma dentata Lutken 1859: 165 (non Muller & Troschel1842); Clark 1921: 121. Location and status of types - MHNG INVE 71892 (Holotype). Type locality- Mauritius. Material examined The specimens collected represent the two known colour forms (Devaney 1970; Sloan et al. 1979). Whole specimens greyish-brown dorsally and ventrally (Plate 2F, G). Disc greyish-brown with fine black reticulating lines, white-ringed black spots, or speckled with light spots. Spots, speckles and lines do not outline the shape of radial shields, thus radial shields are not conspicuous (Plate 3A, B). Oral shields large, round to oval in shape, white on edges The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa and often with large irregular grey blotches. Dental papillae broad but, lower-most shorter than others. Ventral side of arms brown with spotted white bands or with dark bands, with narrower bands in between which continue down the arm. Arm spines taper, annulated white and/or grey. Combination of two and three tentacle scales' on arms. Ecology All specimens were present under large boulders over gravel. Rowe and Gates (1995) record it as a benthic, inshore, littoral species. An overview of known microhabitats of Ophiocoma doederleini is presented in Sloan et al. (1979). Geographical distribution Indian Ocean and west central Pacific Ocean (Rowe & Gates 1995). Our specimens represent a new record for southern Africa. Remarks Devaney (1970: 15-16, Table 2b) provides an accurate means to separate Ophiocoma dentata from 0. doederleini: annulation of the arm spines is easily used in the field and is absent in 0. dentata but present in 0. doederleini. Ophiocoma erinaceus Miiller & Troschel, 1842 (Plate 3C, D) Ophiocoma erinaceus Muller & Troschel 1842: 98; Devaney 1968: 173 (synonymy3); Devaney 1970: 33, figs 45-47; Kalka 1958: 207, 216, 237; Clark 1967: 47; Clark & Rowe 1971:86, 119, pi. 17, figs 5, 6; Clark & Courtman-Stock 1976: 122, 173; Cherbonnier & Guille 1978: 169, pl. X, Figs. 5, 6; Sloan et al. 1979: 106, Figs. 11, 12; Clark 1980: 535, 548; Tortonese 1980: 124; Humphreys 1981: 10, 24; James 1982: 38, pi ID; Price 1982: 8; Clark 1984: 100; Rowe & Gates 1995 : 387; Price & Rowe 1996: 77; Rowe & Richmond 3 Cherbonnier & Guille (1978) removed 0. schoenleinii Miiller & Troschel, 1842 - characterized by the presence of just one tentacle scale, not two as in 0. erinaceus- from Devaney's synonymy. We follow Cherbonnier & Guille (1978). 145 2004: 3292; O'Hara et al. 2004: 537-541; Benavides-Serrato & O'Hara 2008: 51; Reza Fatemi et al. 2010: 44, Fig. 2. Ophiocoma similanensis Bussarawit & Rowe 1985: 1, Figs. 1, 2; Price and Rowe 1996: 77. Location and status of types - ZMB 921 (Syntype 1); ZMB 922 (Syntype 2); ZMB 923 (Syntype 3), specimen lost; ZMB 924 (Syntype 4), specimen lost. Type locality - Red Sea and Indian Ocean. Material examined D.D. 2.8-21.5 mm. Specimens characteristically black, dark brown or dark red dorsally and lighter ventrally. Some of the specimens under study were juveniles as evident from pigmentation of the dorsal disc (pairs of cream lines starting at the margin, passing through the radial shields and almost reaching the centre of the disc; Plate 3D), the arm spines (edge lighter) and the armament of the disc (disc dorsally and ventrally devoid of granules) (see also Cherbonnier & Guille 1978: 171; Bussarawit & Rowe 1985, as Ophiocoma similanensis). Oral shields pearshaped, distally broadest. Dorsal arm plates uniform black, fan-shaped, distally convex, overlapping as tiles on a roof, more than twice as wide as long. Ventral arm plates uniform brown, from regular hexagons proximally to pentagons distally. Two equal-sized tentacle scales over the complete arm. Three to four arm spines, with uppermost one always largest. Three specimens had longitudinal stripes on arms spines similar to the juveniles of this species. Tube feet of the live specimens were reddish. Arm spines on most specimens flattened close to disc. Ecology Benthic, inshore (Rowe & Gates 1995) from 0-27 m depth (Lane et al. 2000). They are associated with coral (Clark & CourtmanStock 1976; Humphreys 1981; Bussarawit & Rowe 1985; StOhr et al. 2008) and are often found on gravel under boulders. Juveniles were found on an encrusting turret sponge (Haliclona sp.) or under dead coral boulders. 146 JENNIFER M. OLBERS and YVES SAMYN An overview of known microhabitats of Ophiocoma erinaceus is presented by Sloan et al. (1979). Geographical distribution Tropical to subtropical Indo-Pacific region. Remarks Even though 0. erinaceus is one of the most abundant and conspicuous brittle stars in littoral tropical seas, its taxonomy has only recently been elaborated. O'Hara et al. (2004) used molecular, morphological and day/night colour change data to show that 0. erinaceus is a species complex of three species: 0. erinaceus, 0. schoenleinii Muller & Troschel, 1842 and a third undescribed species. The latter was formally described and named 0. cynthiae by Benavides-Serrato and O'Hara in 2008. Species in the complex can be distinguished from one another by: (i) colouration of tube feet (red in live or white in preserved 0. erinaceus; grey in life and preserved 0. cynthiae and 0. schoenleinii), (ii) number of tentacle scales (one in 0. schoenleinii; two in 0. erinaceus and 0. cynthiae); (iii) granulation of the ventral side of the disc (largely absent in 0. cynthiae, as a wedge near the margin in 0. schoenleinii and extending almost to the oral shields in 0. erinaceus); and (iv) the size and morphology of the dental plates. Price and Rowe (1996) recognised that their 0. similanensis Bussarawit & Rowe, 1985 is but a juvenile 0. erinaceus and described growth changes for specimens ranging from a D.D. of 3.6-22.2 mm. The juveniles in this study match the description of 0. similanensis very well. Ophiocoma pica Muller & Troschel, 1842 (Plate 3E, F) Ophiocoma pica Muller & Troschel 1842: 101; Clark 1921: 127, pl. 13, Fig. 8; Clark 1938: 333; Balinsky 1957: 25-26; Devaney 1968: 131; Macnae & Kalk 1958: 130; Devaney 1970: 19, Figs. 23 & 24, 20, Figs. 27, 25; Clark & Rowe 1971 : 86-87, 118; Clark & Courtman-Stock 197 6: 173; Cherbonnier & Guille 1978: 172, pl. XI, Figs. 5, 6; Sloan et al. 1979: 106, Clark 1980:535, 548; Tortonese 1980: 124; Price 1982: 8; Clark 1984: 100; James 1982: 36-38, pl IC; Rowe & Gates 1995: 387; Price & Rowe 1996: 77. Location and status of types - Unknown; placed in MNHN according to Muller and Troschel (1842: 101) and according to the MNHN catalogue but not in MNHN according to Dr N. Ameziane (pers. comm.). Type locality -Unknown according to Muller and Troschel (1842). The senior subjective synonym (Lyman 1865) Ophiocoma lineolata Desjardins in Muller and Troschell842 stems from Mauritius. Material examined D.D. = 5.3 mm (D.D./A.L = 1/4). Disc covered dorsally with spherical granules extending onto distal parts of ventral interradii. Colour, pale yellow (after preservation in alcohol). Oral shields mainly oval; adoral shields triangular, not contiguous proximally; oral papillae three to four, dental papillae six to ten; teeth one or two, slightly elongated and blunt. Genital papillae present, cone-shaped. Dorsal arm plates fan-shaped, convex on distal side and concave on proximal side, hardly changing shape distally. Ventral arm plates straight to slightly convex on distal side, concave proximally, plates becoming slightly longer distally. Arm spines five proximally and four distally, slender, second spine longest, about two times segment length, lower arm spines same length as segment or slightly longer. Tentacle scales two, inner one slightly smaller proximally. Ecology Benthic, inshore (Rowe & Gates 1995), 0-24 m (Lane et al. 2000). Usually associated with coral (Clark 1952; Devaney 1968, 1970; Clark & Courtman-Stock 1976; Price & Rowe 1996) but also found under rock or dead coral rubble (Devaney 1970). An overview of known microhabitats of Ophiocoma pica is presented by Sloan et al. (1979). The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa 147 Geographical distribution Widely distributed throughout Indo-Pacific region (Clark 1921; Clark & Rowe 1971). Remarks Given the material from South Africa was in a poor state, we chose to illustrate (Plate 3E, F) a specimen from Mahe (Seychelles). The record discussed here presents a range extension from Richards Bay to Qolora (Eastern Cape Province). Colour in life dark brown or black with radiating golden lines on disc and, often, transverse bands annulating the arms. Ophiocoma pusilla (Brock, 1888) (Plate 4A, B) Ophiomastix pusilla Brock 1888: 499; Devaney 1970: 25 (records before 1970). Ophiocoma latilanxa Murakami 1943a: 194-196; Murakami 1943b: 218; Devaney 1970: 25-27. Ophiocoma pusilla (Brock, 1888); Clark 1921: 131; Devaney 1970: 25, Figs. 26, 29; Cl ark & Rowe 1971: 86-87, 118; Clark & Courtman-Stock 1976: 122, 174, Fig. 190; Cherbonnier & Guille 1978: 173-174, pl. XI, Figs. 3,4; Sloanetal. 1979: 106; Clark 1980: 535, 544; Tortonese 1980: 127; Humphreys 1981: 10, 24; Price 1982: 8; Clark 1984: 100; Rowe & Gates 1995: 388; Price & Rowe 1996: 77. Location and status of types ZMB 5429 (Lectotype); ZMB 4777 (Paralectotype). Type locality Indonesia. Ambon, Plate 4. Dorsal (A) and ventral (B) views of Ophiocoma pusilla; dorsal (C) and ventral (D) views of Ophiocoma scolopendrina (Neotype: MNHN EcOh 11043); dorsal (E) and ventral (F) views of Ophiocoma valenciae. Material examined D.D. = 3.3-7.7 mm. Disc of one specimen slightly speckled while another specimen had banded arms from halfway down the arms to the tips. Dorsal disc with uniformly distributed granules concealing radial shields. Ventral disc with same type of granules, leaving bare only a narrow V-shaped interbrachial area. Oral shields oval, nearly twice as long as wide. Adoral shields triangular, not touching proximally. Four to five oral papillae per jaw. Dental papillae in two to three rows. Dorsal arm plates proximally fan-shaped, wider than long, with convex distal side touching the next plate only for about a third of its width; distally plates longer than wide and less contiguous. Ventral arm plates fan-shaped, broader than long, though distally becoming longer than broad. Four to five arm spines, hollow, glassy and about 2.5 times the segment length. Second uppermost arm spines at a third of arm length JENN1FER M. OLBERS and YVES SAMYN 148 with pustular distal expansions, while other arm spines taper (cf. also Clark & Rowe 1971 ). Tentacle scales, two. Ecology Benthic, inshore (Rowe & Gates 1995), 0-20 m depth according to Lane et al. (2000), our deepest specimen was found at 32 m depth. Known to occur in sand channels, under rubble and associated with coral (Clark & CourtmanStock 1976; Humphreys 1981 ; Price & Rowe 1996). An overview of known microhabitats of Ophiocoma pusilla is presented by Sloan et al. (1979). Geographical distribution Ophiocoma pusilla has a tropical Indo-West Central Pacific Ocean distribution (Rowe & Gates 1995), including the Red Sea (Clark 1967, as Ophiomastix pusilla, Price 1982). In southern Africa, this species was reported from Mozambique (Clark & Courtman-Stock 1976). Remarks After examination of the size, shape and sequence of the arm spines, the nature of the dental plates and oral shields and the number of dental papillae, Devaney (1970) concluded that Ophiocoma latilanxa Murakami, 1943, is a junior synonym of 0. pusilla. In 1989, Soliman ignored this when he identified 0. latilanxa4 from the Red Sea. In 1991, a new species, Ophiocoma aegyptiaca Soliman, 1991, was described from the same area and Soliman (1991) continued to treat 0. latilanxa as a valid species and noted that his new species bears close resemblance to 0. schoenleinii and 0. latilanxa (= 0. pusilla) . Although we have not seen the type material, we suspect that 0. aegyptiaca and 0. latilanxa (= 0. pusilla) will prove to be synonyms. According to Soliman ( 1991 ), the differences between 0. aegyptiaca and 0. latilanxa consist mainly of: (i) the colour pattern of the disc and the plates (our specimens of 0. pusilla show that the colouration is very 4 Soliman 1989 misspelled Ophiocoma latilanxa as 0. latilanaxa. variable (disc uniform brown to reticulate to spotted with dark blotches; spines uniformly coloured to spotted)); (ii) the shape of the oral shields (although Soliman ( 1991) states that the oral shields of 0. aegyptiaca are trapezoidal he drew them oval (Fig. 3A, p. 82 & p. 85), similar to the ones of 0. pusilla); (iii) the number of arm spines (maximum five in 0. aegyptiaca, except Soliman (1991) does not give the sequence of the arm spines; our 0. pusilla specimens mostly have arm spine sequences: 3-3-4-4-4( or 5)-5-4-5). On the other hand the form and size of the dorsal arm plates are very similar for 0. aegyptiaca and 0. pusilla (compare Devaney 1970: Fig. 29 p. 21 with Soliman 1991 : Fig. 4 p. 83). Ophiocoma scolopendrina (Lamarck, 1816) (Plate 4C, D) Ophiura scolopendrina Lamarck, 1816: 544. Ophiocoma scolopendrina (Lamarck, 1816): Kalk 1958: 205 ; Macnae and Kalk 1958: 130; Devaney 1968: 203; Devaney 1970: 33-35; Clark & Rowe 1971: 86, 119, pl. 17, Figs. 3, 4; Clark & Courtrnan-Stock 1976: 122, 174; Sloan et al. 1979: 106, Fig. 13; Clark 1980: 535; Tortonese 1980: 124; Price 1982: 8; James 1982: 36-39, pl. IIA; Rowe & Gates 1995: 388; Reza Fatemi et al. 2010: 45, Fig. 3. Location and status of types - Unknown for 0. scolopendrina, not in the MNHN (Dr M. Eleaume, pers. comm.). According to Devaney (1968), type specimens are present in the Dorpat Museum as the junior subjective synonym 0. variabilis Grube, 1857 (from 'Waohu Island'- Oahu Island?) and in the Museum of Comparative Zoology as the junior subjective synonym 0. molaris Lyman, 1861 (from 'Kingsmill Islands' = Gilbert Islands, Kiribati). Type locality - Recorded by Lamarck (1816) as Mauritius ('lie de France'). The ZMB holds two syntypes of a junior synonym, 0. alternans von Martens, 1870, from Java. The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa Material examined Given that current synonymy might lead to some confusion in its distribution, we felt it appropriate to establish a neotype from the original type locality, Mauritius. Type material N eo type Ophiocoma scolopendrina (Lamarck, 1816), here designated: MNHN EcOh 11043 (specimen with D.D. = 23.8 mm), Mauritius, coil. M. Carie, 1913. Disc uniformly brown both dorsally and ventrally, although where granules have been worn off, lighter patches visible. Dorsal arm plates blotched with brown on beige, giving arms a variegated to banded pattern. Oral and adoral shields with similar blotching. Arm spines uniform in colour (light brown ventrally and somewhat darker dorsally), although on rare occasions some banding can be observed on uppermost spines. Disc pentagonal with interradial margins straight to slightly indented. Dorsal disc densely covered with spherical granules, covering the whole surface including the radial shields which cannot be distinguished. Ventral disc with same, densely distributed granules, but less dense closer to genital slit which is bordered by a fringe of elongated genital papillae. Oral shields oval, shorter (2.5 mm) than wide (3 .0 mm). Adoral shields restricted to the lateral edge of the oral shield, triangular but with other margin curved, not touching proximally. Five oral papillae on each oral plate, inner ones are more pointed than outer, wider ones; buccal tentacle scale very low and wide. Four to nine dental papillae, placed in a cluster below wide truncated teeth. Dorsal arm plates fan-shaped, wider than long with distal margin straight in first segments, becoming more convex in distal segments, plates contiguous throughout the arm. First two ventral arm plates distinctly smaller than rest, with distal margin indented, lateral margins convex and proximal margin straight, about as long as wide. Subsequent ventral arm plates significantly larger, wider then long, with convex distal margin and concave proximal margin which is only slightly overlain by preceding plate, laterally 149 plates recurved around tentacle pore. Arm spines three to five (three on segment three, four to five on segment eight), uppermost ones thick, short, but longer than dorsal arm plates; lower arm spines more slender, always longer than dorsal arm plates, except for first two segments. Arm spines uniform in colour (light brown ventrally and somewhat darker dorsally), although on rare occasion some banding can be observed on uppermost spines. Two oval, tentacle scales, inner one a fraction longer, over the complete arm length. Non-type material To avoid damage to the neotype, we used another specimen to describe the dental plate which is between 1.9 times longer than wide, with a very wide vertical septurn between each oval, elongated tooth foramen; dental papillae region limited to approximately 20% of dental plate length. Remarks As noted by Devaney (1968) and others, 0. scolopendrina is often confused with 0. macroplaca (Clark, 1915), a Hawaiian endemic, and 0. erinaceus Muller & Troschel, 1842. We did not have 0. macroplaca specimens at our disposal so will refrain from commenting on Devaney's (1970) means of distinguishing it from 0. erinaceus and 0. scolopendrina. On the other hand, our limited comparative study of the dental plates of 0. erinaceus, and 0. scolopendrina shows that dental plate morphology can be used to recognize both species with certainty. The senior primary homonym, Ophiocoma scolopendrina Agassiz, 1836 (as for instance mentioned by Muller & Troschel 1842), is to be considered a nomen nudum. Ecology Benthic, inshore, littoral (Rowe & Gates 1995), 0-13 m depth (Lane et al. 2000). An overview of known microhabitats of Ophiocoma scolopendrina is presented by Sloan et al. (1979). 150 JENNIFER M. OLBERS and YVES SAMYN Geographical distribution DISCUSSION Tropical, ludo-Pacific region (Rowe & Gates 1995), including the Red Sea. A total of 70 Ophiocoma specimens belonging to eight species were collected during the five expeditions to KwaZuluNatal. 0. brevipes was the most common (n=24) followed by 0. pusilla (n=l4) and 0. valenciae (n= l4). Only a single individual of 0. dentata was collected in South Africa. Although previously recorded in the Eastern Cape, no 0. pica or 0. scolopendrina were collected by us. 0. brevipes, 0. doederleini, 0. pusilla and 0. dentata are new records for South Africa. Even though Martynov (2010) warned that dental plates are to be used with caution as a taxonomic character, the dental plates permitted us to assign the juveniles in our study to the Brevipes Group, as suggested by Devaney (1970). According to Hendler (1975), only 55 species of ophiuroids have been reported as viviparous, which is less than 3% of all known species. If the juveniles we found attached to adult 0. brevipes individuals are indeed also 0. brevipes individuals, this report is the first account of brooding behaviour and parental care in 0. brevipes. However, based on the examined material, we conclude that freeliving juveniles of 0. brevipes must be very rare, which is in contrast to other ophiocomid species such as 0. erinaceus which have been found as free-living (Price & Rowe 1996). Reproductive experiments with 0. brevipes populations from various locations will probably provide an insight into the ontology and the reproductive strategy of 0. brevipes. Ophiocoma valenciae Muller & Troschel, 1842 (Plate 4E, F) Ophiocoma valenciae Muller & Troschel 1842: 102; Devaney 1968: 126; Eyre & Stephenson 1938: 38, 43; Kalk 1958: 200, 207, 237; Macnae & Kalk 1958: 130; Clark 1967 : 44-45; Macnae & Kalk 1969 : 101, 106, 130; Clark & Rowe 1971: 86, 119, pi. 18 Fig. 1; Sloan et al. 1979: 109, Fig. 14; Clark 1980: 535, 548; Tortonese 1980: 125; Humphreys 1981 : 10, 24-25; Price 1982: 8. Location and status of types - ZMB 4625 (Syntype 1); ZMB 955 (Syntype 2). Type locality - Aden. Material examined D.D. = 7.7-20.3 mm. Disc colour brownish, arms tawny with darker bands; one specimen lacking darker bands on arms. Disc covered dorsally and ventrally with moderately fine granules, which become elongated towards margin of disc. Radial shields defined by lighter colour on some specimens, but could be an artefact due to preservation. Dorsal arm plates broad, oval with up to six arm spines proximally. Uppermost arm spines shorter than middle spines, as long as arm width. Tentacle scale, one. Ecology Associated with coral (Day 1969) and sponges (Humphreys 1981). Found within rocky crevices, cobbles, rubble and various algal beds (Humphreys 1981). An overview of known microhabitats of 0. valenciae is presented by Sloan et al. (1979). Geographical distribution Tropical Indian Ocean, including Red Sea and possibly the Persian Gulf (Clark and Rowe 1971; Tortonese 1980). Acknowledgments-It is with great pleasure that we thank Dr F. Rowe (Senior Research Fellow, Australian Museum, Sydney, Australia) for reading and commenting on the first drafts of this paper. We are also greatly in debt to Dr D. VandenSpiegel who gave us access to collections in the RMCA and who assisted in documenting the specimens. Dr C. Lueter (ZMB), Dr N. Ameziane and Dr M. Eleaume (MNHN), Dr J. Mariaux (MHNG) and Ms S. Halsey (NHM) provided information on the ophiocomid types they curate. We also express our appreciation to Ms K. Vrancken (RBINS) who re-worked some The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa of the figures. The Belgian National Focal Point to the Global Taxonomy Initiative is wannly thanked for financial and logistical support to both authors. The comments of Dr T. O'Hara and an anonymous reviewer greatly improved the paper. REFERENCES AgassizL(l836)Prodromed'unemonographie des radiaires ou Echinodermes. Memoires de la Societe des Sciences Naturelles de Neuchatell: 168-199 Balinsky JB (1957) The Ophiuroidea of Inhaca Island. Annals of the Natal Museum 14: 1-33 Balinsky JB (1969) The Echinoderms. In: Macnae W & Kalk M (eds) Natural History of lnhaca Island: Mozambique. Witwatersrand University Press, Johannesburg, pp 97-156 Bell FJ (1905) Echinoderma found off the coast of South Africa 3. Ophiuroidea. Marine Investigations of South Africa 3: 255-260 Bell FJ (1909) Report on the Echinoderma (other than Holothurians) collected by Mr J. Stanley Gardiner in the Western Parts of the Indian Ocean. Transactions of the Linnaean Society of London: Zoology 13: 17-22 Benavides-Serrato M, O'Hara T (2008) A new species in the Ophiocoma erinaceus complex from the south-west Pacific Ocean (Echinodermata: Ophiuroidea: Ophiocomidae). Memoirs of Museum Victoria 65: 51-56 Brock J (1888) Die Ophiuridenfauna des indischen Archipels. Zeitschrift flir Wissenschaftliche Zoofogie 47: 465-539 Bussarawit S, Rowe FWE (1985) A new species in the Ophiocomid genus Ophiocoma (Echinodermata: Ophiuroidea) from the west coast of Thailand, Andaman Sea. Research Bulletin Phuket Marine Biological Centre, Research Bulletin 35: 1-6 151 Cherbonnier G, Guille A (1978) Faune de Madagascar 48. Echinodermes: Ophiurides. Museum National de' Historie Naturelle, 272 pp Clark AH (1952) Echinoderms from the Marshall Islands. Proceedings of the United States National Museum 102(3302): 265-303 Clark AM (1967) Echinoderms from the Red Sea. Part 2. (Crinoids, Ophiuroids, Echinoids and more Asteroids). Bulletin of the Sea Fisheries Research Station Israel41: 26-58 ClarkAM ( 197 4) Notes on some echinoderms from South Africa. Bulletin of the British Museum of Natural History (Zoology) 26 : 421-487 Clark AM (1977) The South African Museum's Meiring Naude cruises. 4. Echinoderms. Annals of the South African Museum 76: 13-147 Clark AM (1980) Some Ophiuroidea from the Seychelles Islands and lnhaca, Mozambique. Revue de Zoologie Africaine 94: 533-558 Clark AM (1984) Echinodermata of the Seychelles. In D. R. Stoddart (ed) Biogeography and ecology of the Seychelles Islands. Dr W Junk Publishers, The Hague, pp 83-102 Clark AM, Rowe FWE (1971) Monograph of shallow-water Indo-West Pacific Echinoderms. Publication no. 690. British Museum of Natural History, London, 23 8 pp Clark AM, Courtman-Stock J (1976) The Echinoderms of Southern Africa. British Museum (Natural History), London, 277 pp Clark HL (1908) Some Japanese and East Indian echinoderms. Bulletin of the Museum of Comparative Zoology 11(11): 279-311 Clark HL (1911) North Pacific Ophiurans in the collection of the United States National Museum. Bulletin of the United States National Museum 75, 336 pp ..... 152 JENNIFER M. OLBERS and YVES SAMYN Clark HL (1915) Catalogue of recent Ophiurans, based on the collection of the Museum of Comparative Zoology. Memoirs of the Museum of Comparative Zoology at Harvard College 25 : 164-376 Eyre J, Stephenson TA (1938) The South African intertidal zone and its relation to ocean currents. V. A subtropical Indian Ocean shore. Annals of the Natal Museum 9: 21-46 Clark HL (1921) The echinoderm fauna of Torres Strait. Publications of the Carnegie Institute Washington 10: 1-233 Fourgon D, Jangoux M, Eeckhaut I (2007) Biology of a 'babysitting' symbiosis in brittle stars: analysis of the interactions between Ophiomastix venosa and Ophiocoma scolopendrina. Invertebrate Biology 126: 385-395 Clark HL (1923) The Echinoderm fauna of South Africa. Annals of the South African Museum 13 : 221-435 Clark HL (1938) Echinoderms from Australia, an account of collections made in 1929 and 1932. Memoirs of the Museum of Comparative Zoology at Harvard College 55: 727 Day lli (1969) A guide to marine life on South African shores. Cape Town, Balkema, 300 pp de Loriol P (1893) Catalogue raisonne des Echinodermes recueillis par M. V. de Robillard a 1'lie Maurice. Ill. Ophiurides et Astrophytides. Memoires de la Societe de Physique et d'Historie Naturelle de Geneve 32: 1-64 de Loriol P (1899) Notes pour server a !'etude des echinoderms. VII. Memoires de la Societe de Physique et d 'Historie Naturelle de Geneve 33(2, 1): 1-34 Devaney DM (1968) The systematics and post-larval growth changes in Ophiocomid brittle stars. PhD thesis. University of Hawaii, 272 pp Devaney DM (1970) Studies on Ophiocomid brittle stars.!. Anew Genus (Clarkcoma) of Ophiocominae with a re-evaluation of the Genus Ophiocoma. Smithsonian Contributions to Zoology 51: 1-41 Devaney DM (1974) Shallow-water asterozoans of southeastern Polynesia. 2. Ophiuroidea. Micronesica 10: 105-204 Devaney DM (1978) A review of the Ophiomastix (Ophiuroidea: genus Ophiocomidae). Micronesica 14: 273-359 Grube AE (1857) Diagnosen einiger neuen Echinodermen. Archiv Ffu Naturgeschicthe 23: 340-344 Hendler G (1975) Adaptational significance ofthe patterns ofophiuroid development. American Zoologist 15: 691-715 Hendler G (1984) Brittlestar colour-change and phototaxis (Echinodermata: Ophiuroidea: Ophiocomidae) . Marine Ecology 5: 379-401 Hyman LH (1955) The invertebrates: Echinodermata, The Coelomate Bilateria. New York, McGraw-Hill Book Company, 763 pp James DB (1982) Studies on Indian echinoderms-1 0. Ophiocomaanaglyptica (Ophiuroidea: Ophiocomidae), a new record from the Indian Ocean with notes on other species of Ophiocoma from Indian seas. Journal of the Marine Biological Association oflndia 24: 36-41 Kalk M (1958) The fauna of the intertidal rocks at Inhaca Island, Delagoa Bay. Annals of the Natal Museum 14: 189-242 Koehler R (1922) Ophiurians of the Philippine seas and adjacent waters. Bulletin of the United States National Museum 100: 1-480 Lamarck, de JBPA (1816) Historie Naturelle des Animaux sans Vertebres. Verdiere, Paris 2: 522-568 The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa Lane DJW, Marsh LM, VandenSpiegel D, Rowe FWE (2000) Echinoderm fauna of the South China Sea: An inventory and analysis of distribution patterns. The Raffles Bulletin of Zoology 8(supplement): 459-493 Ljungman A (1867) Om nagra nya arter af Ophiurider. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar, pp 163-166 Llitken C (1859) Bidrag til Kundskab om de ved Kysteme af Mellem og SydAmerika 1evende Arter af Sostjemer. Videnskabelige Meddelelser, pp 25-96 Lyman T (1861) Descriptions of new Ophiuridae, belonging to the Smithsonian Institution and to the Museum of Comparative Zoology at Cambridge. Proceedings of the Boston Society ofNatural History 7: 193-203 153 Matsumoto H (1915) A new classification of the Ophiuroidea: with descriptions of new genera and species. Proceedings of the Academy of Natural Sciences of Philadelphia 67 : 43-92 Mortensen T (1925) On some echinoderms from South Africa. Annals and Magazine of Natural History 9: 146-154 Mortensen T (1933) Echinoderms of South Africa (Asteroidea and Ophiuroidea). Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 93: 215-490 Muller J, Troschel FH (1840) Ober die Gattungen der Ophiuren. Wiegmanns Archiv fur Naturgeschichte 6: 318-326 Muller J, Troschel FH (1842) System der Asteriden. Braunschweig, 134 pp Lyman T (1862) Descriptions of new Ophiuridae. Proceedings of the Boston Society ofNatural History 8: 75-86 Murakami S ( 1943a) Report on the Ophiurans of Palao Caroline Islands. Journal of the Department of Agriculture, Kyusyu Imperial University 7: 159-203 Lyman T (1865) Ophiuridae and Astrophytidae. Illustrated Catalogue of the Museum of Comparative Zoology, Harvard University 1: 1-200 M urakami S ( 1943b) Report on the Ophiurans of Yaeyama, Ryukyu. Journal of the Department of Agriculture, Kyusyu Imperial University 7: 205-222 Macnae W, Kalk M (1958) A natural history of lnhaca Island, Mozambique. Witwatersrand University Press, Johannesburg, 163 pp O'Hara TD, Byre M, Cistemas P (2004) The Ophiocoma erinaceus complex: another case of cryptic speciation in echinoderms: pp 537-542 In: Heinzeller T, Nebelsick JH (eds) Echinoderms Munchen: Proceedings of the 11th International Echinoderm Conference, Munich, Germany, 6-10 October 2003, A.A. Balkema, Netherlands. Macnae W, Kalk M ( 1969) A natural history of Inhaca Island, Mozambique (Second edition). Witwatersrand University Press, Johannesburg, 395 pp (1887) G Marktanner-Turneretscher Beschreibung neuer Ophiuriden und Bemerkungen zu Bekannten. Annalen des Naturhistorischen · Museums in Wien 2:291-316 Martynov, A (2010) Reassessment of the classification of the Ophiuroidea (Echinodermata), based on morphological characters. I. General character evaluation and delineation of the families Ophiomyxidae and Ophiacanthidae. Zootaxa 2697: 1-154 Peters WCH (1851) Ubersicht der von ihm an der Kuste von Mossambique eingesammelten Ophiuren, unter denen sich zwei neue Gattungen befinden. Berichte der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin 1851 : 463-466 154 JENNIFER M. OLBERS and YVES SAMYN Price ARG, Rowe FWE (1996) Indian Ocean echinoderms collected during the Sinbad Voyage (1980-1981): 3 Ophiuroidea and Echinoidea. Bulletin of the Natural History Museum ofLondon (Zoology) 62: 71-82 Price ARG (1982) Echinoderms of Saudi Arabia. Comparison between echinoderm faunas of Arabian Gulf Sea, SE Arabia, Red Sea, Gulfs of Aqaba and Suez. Fauna Saudi Arabia 4: 3-21 Reza Fatemi SM, Jamili S, Valinassab T, Kuranlu N (2010) Diversity of Ophiuroidea from Lengeh Portand Qeshm Island in the Persian Gulf. Journal of Fisheries and Aquatic Science 5: 42-48 Rowe FWE, Gates J (1995) Echinodermata. In: Wells A (ed.) Zoological Catalogue of Australia 33, Melbourne: CSIRO Australia viii, 510 pp Rowe FWE, Pawson DL (1977) A catalogue of echinoderm type-specimens in the Australian Museum, Sydney. Records of the Australian Museum 30: 337-364 Rowe FWE, Richmond MD (2004) A preliminary account of the shallow water echinoderms of Rodrigues, Mauritius, Western Indian Ocean. Journal of Natural History 38: 3273-3314 Samyn Y, Thandar AS (2003). Towards an understanding of the shallow-water echinoderm biodiversity of KwaZuluNatal, Republic of South Africa. Echinoderm Research 2001: 41-4 7 Sloan NA, Clark AM, Taylor JD (1979) . The echinoderms of Aldabra and their habitats. Bulletin of the British Museum (Natural History), Zoology 37: 81-128 Soliman FE (1989) Studies on the Egyptian Echinodermata. Ophiocoma latilanxa (Ophiuroidea: Ophiocomidae). A new record from the Red Sea. Bulletin of the Faculty of Science of Assiut University 18: 97-105 Soliman FE (1991) Studies on Egyptian Echinodermata: Ophiocoma aegyptiaca sp. nov. (Ophiuroidea: Ophiocomidae) from the Red Sea. Galaxea 10: 79-88 St6hr S (2005) Who 's who among baby brittle stars (Echinodermata: Ophiuroidea) : post metamorphic development of some North Atlantic forms. Zoological Journal of the Linnaean Society 143: 543-576 St6hr, S, Conand C, Boissin E (2008) Brittle stars (Echinodermata: Ophiuroidea) from La Reunion and the systematic position of Ophiocanops Koehler 1922. Zoological Journal of the Linnaean Society 153: 545-560 Surnida PYG, Tyler PA, Gage JD, Norrevang A (1998) Postlarval development in shallow and deep-sea ophiuroids (Echinodermata: Ophiuroidea) of the NE Atlantic Ocean. Zoological Journal of the Linnaean Society 124: 267-300 Tortonese E (1980) Researches on the coast of Somalia. Littoral Echinodermata, Monitore Zoologico ltaliano N . S. Supplemento 13, 5: 99- 139. von Martens E (1870) Die Ophiuriden des indischen Oceans. Archiv fur Naturgeschichte 36: 244-262.